FB2026_02 , released June 18, 2026
Gene: Dmel\Tlk
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General Information
Symbol
Dmel\Tlk
Species
D. melanogaster
Name
Tousled-like kinase
Annotation Symbol
CG34412
Feature Type
FlyBase ID
FBgn0283657
Gene Model Status
Stock Availability
Gene Summary
Tousled-like kinase (Tlk) encodes a serine/threonine kinase that acts in a cell cycle-dependent manner. It contributes to cell morphogenesis, cell migration and cell death through interactions with chromatin components. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)
Also Known As

l(1)G0054, EP1413, BEST:GH17330 , BcDNA:GH07910

Key Links
Genomic Location
Cytogenetic map
Sequence location
Recombination map
1-4
RefSeq locus
NC_004354 REGION:3720007..3789770
Sequence
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
Gene Ontology (GO) Annotations (11 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
inferred from physical interaction with FLYBASE:asf1; FB:FBgn0029094
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
Biological Process (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
inferred from genetic interaction with FLYBASE:asf1; FB:FBgn0029094
inferred from genetic interaction with FLYBASE:mars; FB:FBgn0033845
inferred from mutant phenotype
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
inferred from biological aspect of ancestor with PANTHER:PTN000540737
inferred from biological aspect of ancestor with PANTHER:PTN000540738
Cellular Component (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
located_in heterochromatin
inferred from direct assay
located_in nucleoplasm
inferred from high throughput direct assay
located_in nucleus
inferred from direct assay
Terms Based on Predictions or Assertions (0 terms)
Protein Family (UniProt)
-
Summaries
Gene Snapshot
Tousled-like kinase (Tlk) encodes a serine/threonine kinase that acts in a cell cycle-dependent manner. It contributes to cell morphogenesis, cell migration and cell death through interactions with chromatin components. [Date last reviewed: 2019-03-14]
Gene Group (FlyBase)
UNCLASSIFIED CONVENTIONAL PROTEIN KINASE DOMAIN KINASES -
This group comprises conventional protein kinase domain kinases that do not classify under other groups in FlyBase.
Gene Model and Products
Number of Transcripts
14
Number of Unique Polypeptides
10

Please see the JBrowse view of Dmel\Tlk for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry B7Z126)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Gene model reviewed during 5.55

3' UTR contains an ORF that is conserved among close Drosophila species (FBrf0202906; potential peptide 129 aa in length).

Gene model reviewed during 6.46

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated

Due to significant sequence change in this region between Releases 4 and 5, data that predate Release 5 are not shown for some categories of evidence.

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.46

Stop-codon suppression (UAG) postulated; FBrf0243886.

Gene model reviewed during 6.32

Double stop-codon suppression (UAG, UAA) postulated; FBrf0243886.

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0112680
4624
1463
FBtr0070596
5849
1254
FBtr0299579
5537
1255
FBtr0299580
6513
1266
FBtr0299581
7243
1489
FBtr0299582
5890
1266
FBtr0299583
4673
1242
FBtr0301659
4882
1266
FBtr0333751
7403
1265
FBtr0333752
4308
1011
FBtr0333753
7490
1242
FBtr0345319
4355
1242
FBtr0479803
5890
1276
FBtr0481732
5890
1412
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
FBpp0111592
160.3
1463
9.10
FBpp0099673
135.5
1254
9.31
FBpp0288854
135.6
1255
9.36
FBpp0288855
136.7
1266
9.10
FBpp0288856
162.2
1489
9.62
FBpp0288857
136.7
1266
9.10
FBpp0288858
133.9
1242
9.02
FBpp0290873
136.7
1266
9.10
FBpp0305893
136.4
1265
9.25
FBpp0305894
110.0
1011
9.50
FBpp0305895
133.9
1242
9.02
FBpp0311479
133.9
1242
9.02
FBpp0428136
137.8
1276
9.24
FBpp0428486
152.7
1412
9.20
Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

1266 aa isoforms: Tlk-PF, Tlk-PH, Tlk-PJ
1242 aa isoforms: Tlk-PI, Tlk-PM, Tlk-PN
Additional Polypeptide Data and Comments
Reported size (kDa)
Comments
External Data
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\Tlk using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

-0.77

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
nurse cell & oocyte

Comment: maternal transcript transported from nurse cells to oocytes

northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

tlk is ubiquituously distributed in all proliferative tissues.

Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data

tlk protein is localized to the nucleus in interphase cells, and diffusely distributed throughout the cell during other parts of the mitotic cell cycle. tlk protein does not associate directly with DNA.

Marker for
 
Subcellular Localization
CV Term
Evidence
References
located_in heterochromatin
inferred from direct assay
located_in nucleoplasm
inferred from high throughput direct assay
located_in nucleus
inferred from direct assay
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\Tlk in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Images
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 88 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 22 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Tlk
Transgenic constructs containing regulatory region of Tlk
Aberrations (Deficiencies and Duplications) ( 0 )
Inferred from experimentation ( 0 )
Inferred from location ( 4 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Sterility
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
larval salivary gland & nuclear chromosome, with Scer\GAL4hs.2sev
Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (15)
10 of 14
Yes
Yes
9 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
5  
1 of 14
No
No
4  
1 of 14
No
No
2  
1 of 14
No
No
4  
1 of 14
No
No
2  
1 of 14
No
No
2  
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1  
1 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (10)
10 of 14
Yes
Yes
8 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Mus musculus (laboratory mouse) (16)
10 of 14
Yes
Yes
9 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (16)
7 of 13
Yes
Yes
5 of 13
No
No
4 of 13
No
Yes
4 of 13
No
Yes
4 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
Danio rerio (Zebrafish) (13)
10 of 14
Yes
Yes
8 of 14
No
Yes
8 of 14
No
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (4)
11 of 14
Yes
Yes
1 of 14
No
No
1 of 14
No
No
1 of 14
No
No
Anopheles gambiae (African malaria mosquito) (6)
9 of 12
Yes
Yes
Arabidopsis thaliana (thale-cress) (17)
9 of 13
Yes
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
No
1 of 13
No
No
1 of 13
No
No
1 of 13
No
Yes
1 of 13
No
Yes
Saccharomyces cerevisiae (Brewer's yeast) (11)
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
Schizosaccharomyces pombe (Fission yeast) (6)
1 of 12
Yes
No
1 of 12
Yes
No
1 of 12
Yes
No
1 of 12
Yes
No
1 of 12
Yes
No
1 of 12
Yes
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:Tlk. Refer to their site for version information.
Paralogs
Paralogs (via DIOPT v9.1)
Drosophila melanogaster (Fruit fly) (12)
2 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
1 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 1 )
    Modifiers Based on Experimental Evidence ( 1 )
    Allele
    Disease
    Interaction
    References
    Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Functional Complementation Data
    Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
    Interactions
    Summary of Physical Interactions
    Summary of Genetic Interactions
    Interaction Browsers

    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    External Data
    Linkouts
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Signaling Pathways (FlyBase)
    Metabolic Pathways
    FlyBase
    External Links
    External Data
    Linkouts
    Class of Gene
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    X
    Recombination map
    1-4
    Cytogenetic map
    Sequence location
    FlyBase Computed Cytological Location
    Cytogenetic map
    Evidence for location
    3E6-3F1
    Left limit from in situ hybridisation (FBrf0128367) Right limit from in situ hybridisation (FBrf0128367); Limits computationally determined from genome sequence between P{EP}EP1200EP1200&P{EP}EP1343 and P{EP}CG2930EP1352 3F1--5
    Experimentally Determined Cytological Location
    Cytogenetic map
    Notes
    References
    3F4-3F5
    (determined by in situ hybridisation)
    3E6-3E7
    (determined by in situ hybridisation)
    3F1-3F5
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Location
    Left of (cM)
    Right of (cM)
    Notes
    Stocks and Reagents
    Stocks (56)
    Genomic Clones (26)
    cDNA Clones (101)
     

    Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see JBrowse for alignment of the cDNAs and ESTs to the gene model.

    cDNA clones, fully sequenced
    Other clones
      Drosophila Genomics Resource Center cDNA clones

      For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

      cDNA Clones, End Sequenced (ESTs)
      RNAi and Array Information
      Linkouts
      Antibody Information
      Laboratory Generated Antibodies
       

      polyclonal

      Commercially Available Antibodies
       
      Cell Line Information
      Publicly Available Cell Lines
       
        Other Stable Cell Lines
         
          Other Comments

          RNAi screen using dsRNA made from templates generated with primers directed against this gene causes decreased γ-tubulin staining at the spindle pole and misalignment of chromosomes along the spindle when assayed in S2 cells. This phenotype can be observed when the screen is performed with or without Cdc27 dsRNA.

          Annotation CG12462 eliminated in release 5.1 of the genome annotation.

          RNAi screen using dsRNA made from templates generated with primers directed against this gene causes a greater than three-fold increase in AttA activity in response to heat-killed E.coli after ecdysone treatment in S2 cells.

          When dsRNA constructs are made and transiently transfected into S2 cells in RNAi experiments, an increase in the proportion of S phase cells, G2/M+, an increase in the proportion of G2/M phase cells, a decrease in cytokinetic index and a decrease in the mitotic index are seen.

          New annotation (CG33219) in release 3.2 of the genome annotation.

          The release 4 CG32782 annotation contains sequences corresponding to release 2 annotation CG2829. The release 3 version of CG32782 was incomplete and was missing most of the sequence corresponding to release 2 annotation CG2829 (possibly due to an assembly error), but this has been fixed in release 4 and the release 4 CG32782 annotation now includes all of the release 2 CG2829 sequence. 'CG33219' does not appear in Release 4. The region it was located in in Release 3.2 is now a large TE. 'CG33219' is homologous to annotation CG32782; presumably that is where this little bit of mislaid 'CG33219' sequence originated.

          The tlk-asf1 pathway is important for both DNA replication and chromatin assembly. tlk may coordinate cell cycle progression through the regulation of chromatin dynamics.

          dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology.

          RNAi screen using dsRNA made from templates generated with primers directed against this gene causes a phenotype when assayed in Kc167 and S2R+ cells: cells become round and detached.

          New annotation (CG32782) in release 3 of the genome annotation. Annotation CG2829 was eliminated in release 3 of the genome annotation, October 2002.

          New annotation (CG32781) in release 3 of the genome annotation.

          Relationship to Other Genes
          Source for database merge of

          Source for merge of: tlk anon-WO03040301.114 anon-WO03040301.116

          Source for merge of: tlk CG32781

          Source for merge of: CG12462 EP1413

          Source for merge of: CG32782 CG33219

          Source for merge of: tlk CG12462

          Source for merge of: tlk CG32782

          Source for merge of: Tlk EP1408

          Additional comments

          Annotations CG32782 and CG32781 merged as CG34412 in release 5.2 of the genome annotation.

          Sequences corresponding to annotation CG12462 (which was eliminated in release 5.1 of the genome annotation) have been reinstated as part of the CG34412 annotation (which corresponds to tlk) in release 5.2 of the genome annotation.

          "l(1)G0299" may affect "CG2829". "l(1)G0167" may affect "CG2829". "l(1)G0343" may affect "CG2829". "l(1)G0054" may affect "CG2829".

          Source for merge of tlk anon-WO03040301.114 anon-WO03040301.116 was sequence comparison ( date:051113 ).

          Source for merge of tlk CG32782 was sequence comparison ( date:050324 ).

          Nomenclature History
          Source for database identify of

          Source for identity of: Tlk tlk

          Nomenclature comments
          Etymology
          Synonyms and Secondary IDs (23)
          Reported As
          Symbol Synonym
          BEST:GH17330
          BcDNA:GH07910
          EP1408
          anon-WO03040301.114
          anon-WO03040301.116
          Secondary FlyBase IDs
          • FBgn0086899
          • FBgn0052781
          • FBan0032782
          • FBgn0085441
          • FBgn0053219
          • FBgn0052782
          • FBgn0066856
          • FBgn0066855
          • FBgn0028489
          • FBgn0026698
          • FBgn0086709
          • FBgn0062688
          • FBgn0029675
          • FBgn0043460
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          External Crossreferences and Linkouts ( 79 )
          Sequence Crossreferences
          NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
          GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
          GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
          UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
          UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
          Other crossreferences
          AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
          BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
          DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
          EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
          FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
          KEGG Genes - Molecular building blocks of life in the genomic space.
          MARRVEL_MODEL - MARRVEL (model organism gene)
          Linkouts
          Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
          Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
          FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
          FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
          Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
          iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
          MIST (genetic) - An integrated Molecular Interaction Database
          MIST (protein-protein) - An integrated Molecular Interaction Database
          References (145)